The Mantispidae (Insecta: Neuroptera) of Canada, with Notes on Morphology, Ecology, and Distribution1

531 The Mantispidae (Insecta: Neuroptera) of Canada, with notes on morphology, ecology, and distribution1

Robert A. Cannings2 Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia, Canada V8W 9W2 Sydney G. Cannings NatureServe Yukon, Yukon Territorial Government, Box 2703, Whitehorse, Yukon Territories, Canada Y1A 2C6, and Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia, Canada V8W 9W2

Abstract—Morphological, biological, and distributional data are presented for the four Cana- dian species of Mantispidae. A key for the identification of these species is provided. Climaciella brunnea (Say) is the most frequently collected Canadian species, occurring in southern British Columbia, Alberta, Manitoba, Ontario, and Quebec. Dicromantispa interrupta (Say) comb. nov. is given as a new combination for Mantispa interrupta Say; it is recorded in southern Ontario and Quebec. Dicromantispa sayi (Banks) occurs only in extreme southwestern Ontario. Leptomantispa pulchella (Banks) inhabits the dry Okanagan Valley of British Columbia; it is also known from Ojibway Prairie, Windsor, Ontario. The presence of the latter two species in Canada is published for the first time.

Résumé—Nous présentons des données sur la morphologie, la biologie et la répartition des quatre espèces canadiennes de Mantispidae, ainsi qu’une clef pour leur identification. Clima- ciella brunnea (Say) est l’espèce canadienne la plus fréquemment récoltée et elle se retrouve dans le sud de la Colombie-Britannique, de l’Alberta, du Manitoba, de l’Ontario et du Québec. Dicromantispa interrupta (Say) comb. nov. est une nouvelle combinaison pour désigner Man- tispa interrupta Say; l’espèce habite le sud du Québec et de l’Ontario. Dicromantispa sayi (Banks) se rencontre seulement dans l’extrême sud-ouest de l’Ontario. Leptomantispa pulchella (Banks) se retrouve dans la vallée sèche de l’Okanagan en Colombie-Britannique; elle a été ré- coltée aussi à Ojibway Prairie, Windsor, Ontario. La présence de ces deux dernières espèces au Canada est signalée pour la première fois.

[Traduit par la Rédaction]

Cannings544 and Cannings Introduction being that the Mantispidae are holometabolous and the Mantidae are not. Adults of the neuropteran family Manti- spidae, with their triangular head and large The four species of Mantispidae known eyes, elongate prothorax and raptorial forelegs, from Canada are Climaciella brunnea (Say), strikingly resemble the Mantidae of the order Dicromantispa interrupta (Say) comb. nov., Mantodea. However, the two families are only Dicromantispa sayi (Banks), and Leptoman- distantly related in the Insecta; the superficial re- tispa pulchella (Banks). Until recently, the last semblance is the result of convergent evolution. three species had been placed in Mantispa,but There are many differences, the most obvious Hoffman (2002) created new genera for the

Received 20 June 2004. Accepted 16 September 2005. 1This paper is part of a special issue honouring Geoffrey G.E. Scudder for his significant contributions to entomology in Canada. 2Corresponding author (e-mail: [email protected]).

Nearctic and Neotropical species of the In the Mantispinae, first-instar larvae are subfamily Mantispidae based on his findings campodeiform and active and find spider eggs that Old and New World clades were distinct by searching for, and penetrating, egg sacs or (Hoffman 1992; K. Hoffman, personal commu- by boarding a female spider and entering the nication). egg sac as it is constructed (Redborg and Species of Mantispidae in Canada are at the MacLeod 1985). In some North American man- northern limits of their range; none are recorded tispines studied, one of these two behaviours is north of 49°N latitude in Ontario and Quebec obligatory, although in D. sayi, either is possi- and none north of 51°N latitude west of Ontario. ble. This is probably also the case in D. inter- However, this fauna represents the majority of rupta (Redborg 1998). Larvae of C. brunnea Nearctic mantispines, because the most recent and L. pulchella are apparently obligate spider taxonomic work indicates that only six species boarders. Larvae often feed on spider haemo- of Mantispinae occur in North America: the four lymph while riding on the host (Redborg and species recorded in Canada plus Xeromantispa MacLeod 1984). Zeugomantispa minuta scabrosa (Banks, 1912) and Zeugomantispa (= Mantispa viridis Walker) actively searches minuta (Fabr., 1775) (K. Hoffman, personal out egg sacs and chews its way into them communication). This fauna is an extension of a (Redborg and MacLeod 1985) and will feed on primarily Central and South American one: all eggs presented to it (Davidson 1969). Accord- six species occur in Mexico and all except ing to much of the literature, second- and third- D. interrupta and X. scabrosa range at least as instar larvae become sedentary and eruciform far south as Nicaragua (Hoffman 1992). inside the egg cases (Imms 1957), although Canadian mantispids belong to the subfamily Redborg (1998) disputed the idea that mantispids Mantispinae, apparently among the more de- are truly hypermetamorphic in the way, for ex- rived mantispids, whose members all develop, ample, meloid beetles are. The entire food sup- as far as is known, by feeding on spider eggs in ply for the insect’s development is inside a single the larval stages. Other mantispids are less well egg sac; adult size varies considerably and de- known biologically and are considered more pends on the amount of food eaten by the larva plesiotypic, the larvae, at least in some cases, (Redborg and MacLeod 1985). After pupation, remaining generalist predators (MacLeod and the pharate adult chews its way out of the pupal Redborg 1982; Redborg 1998). cocoon and the egg sac before final adult ecdysis.

Key to the Canadian species of Mantispidae 1a Wings with anterior half amber to brown; in anterior view, antennal flagellomeres at midlength of flagellum each 3 or more times as wide as long; pronotum with strong constrictions and with transverse ridges; pronotum in dorsal view usually 4 to 5.5 times as long as wide at the narrowest point; one species widely distributed in southern Canada from Quebec west to British Columbia . . . Climaciella brunnea 1b Wings mostly clear, with any brown markings restricted to the costal/radial area; in anterior view, antennal flagellomeres at midlength of flagellum each less than 3 times as wide as long; pronotum without strong constrictions and with or without transverse ridges; pronotum in dorsal view usually 6 or more times as long as wide at the narrowest point ...... 2 2a Pronotum in lateral view with numerous short setae over its entire length; pronotum without evident transverse ridges; a species of extreme southwestern Ontario and dry valleys in the southern interior of British Columbia ...... Leptomantispa pulchella 2b Pronotum in lateral view with scattered fine setae, most of them at the anterior and posterior ends of the pronotum; pronotum with evident transverse ridges; species from Ontario and Quebec ...... 3 3a Wing tips and some crossveins of radial cells marked with brown spots . . . Dicromantispa interrupta 3b Wing tips and crossveins without brown spots ...... Dicromantispa sayi

Specimens cited in the species accounts are BCAK British Columbia Ministry of Agri- housed in the following collections (abbrevia- culture and Lands, Kelowna, British tions follow Arnett et al. (1993)): Columbia, Canada

BCPM Royal British Columbia Museum, Mantispa burquei Provancher, 1875. Victoria, British Columbia, Canada Climaciella brunnea (Say), Enderlein (1910); BDMU McMaster University, Hamilton, On- synonymy of M. burquei; new genus. tario, Canada Climaciella brunnea (Say) occidentis Banks, BMNH Natural History Museum, London, 1911; description of western form. United Kingdom CNCI Canadian National Collection of In- Range sects, Arachnids and Nematodes, Southern British Columbia and Alberta Agriculture and Agri-Food Canada, (probably occurs in southern Saskatchewan, but Ottawa, Ontario, Canada no specimens known), Manitoba, Ontario, and DEBU University of Guelph, Guelph, On- Quebec south to California, Arizona, New Mex- tario, Canada ico, Mississippi, and Florida; south through DNC Dean Nicholson Collection, Cran- Mexico to Costa Rica (see Throne (1972) for a brook, British Columbia, Canada summary of the distribution in the United EDUM University of Manitoba, Winnipeg, States). Manitoba, Canada GLFR Great Lakes Forest Research Centre, Canadian specimens examined Sault Ste. Marie, Ontario, Canada British Columbia: Craig, 2.vii.1953, Forest IFE S. Ife Collection Insect Survey (PFRS); Cranbrook, 3400 feet, LEMQ Lyman Entomological Museum, 5.vii.2004, J. Collings (DNC); Goldstream, McGill University, Sainte-Anne-de- 9.vi.1917, E.H. Blackmore (CNCI); Kamloops, Bellevue, Quebec, Canada 9.vii.1944, 3.ix.1944, G.J. Spencer (SMDV); LUTB Lakehead University, Thunder Bay, Kelowna, 8.vii.1968, Forest Insect Survey Ontario, Canada (CNCI); Kootenay Valley, 2500′, 1900–1922, MCZC Museum of Comparative Zoology, C.H. Polson (BMNH); Lillooet, 25.ix.1918, Harvard University, Cambridge, A.W.A. Phair (CNCI); Marron Lake, 26.ix.1923, Massachusetts, United States of C. de B. Green (CNCI); vii.1924 (3 specimens), America C. de B. Green (BCPM); Okanagan Falls, NFRC Northern Forest Research Centre, Willowbrook, 12.viii.2002, M. Sarell (BCPM); Edmonton, Alberta, Canada Okanagan Landing, vi.1990, H. Philip (SMDV); OSUC Ohio State University, Columbus, Oliver, 25.viii.1950, P. Zuk (SMDV); Ohio, United States of America 20.viii.1975, P.J. Procter (BCPM); ix.1991, T. PFRS Pacific Forestry Centre, Victoria, Watson (BCAK); 11.ix.2000, D. Nemeth British Columbia, Canada (SMDV); Oliver, 5 mi. N, 6.ix.1962, M. Preusse QMOR Université de Montréal, Montréal, (SMDV); Osoyoos, no date, C. de B. Green Quebec, Canada (CNCI); 8.ix.1983, J. Belton (BCPM); Oyama, ROME Royal Ontario Museum, Toronto, 20.ix.1926, J.A. Henderson (BCPM); Ontario, Canada 20.x.1929, G.J. Spencer (SMDV); Roosville, SMDV Spencer Entomological Museum, Tobacco Plains, 17.vii.1999, M. Fairbarns and University of British Columbia, Van- S. Ford (BCPM); Summerland, 27.viii.1920, couver, British Columbia, Canada F.W. Sladen (CNCI); 9.ix.1981, P.J. Procter USNM United States National Museum, (BCAK); Trepanier, ix.1943, V. Nelson Washington, District of Columbia, (SMDV); Vaseux Lake, 26.vi.1959, L.A. Kelton United States of America (CNCI), 20.viii.1989 (3 specimens), S. Ife UVIC University of Victoria, Victoria, (IFE); Vernon, 5.x.1940, G.R.N. (SMDV); British Columbia, Canada Vernon, Commonage, 12.vi.2005, K. Iverson (BCPM); Vernon, Cosens Bay, 21.vii.1987, R.A. Cannings (BCPM); Victoria, no date, Rev. Species accounts G.W. Taylor (MCZC), 1917 (USNM). Manitoba: Beaver Creek, 26.vi.1962, J.A. Climaciella brunnea (Say, 1824) Garland (EDUM); Birdshill Provincial Park, 25.vi.1978, D.G. Delf (EDUM); Fraserwood, Mantispa brunnea Say, 1824; original 21.vii.1964, Forest Insect Survey (CNCI); description; syntype, St. Peter’s River, Lockport, 28.vi.1976, 7.vii.1976, J.S. Lee Minnesota, probably destroyed. (EDUM); St. Laurent, 23.vii.1961, Forest Insect

Survey W61–1672–02 (NFRC); Tenlon, 24.vi.1975, M. Sanborne (LEMQ); Pelee Is- 4.vii.1958, M. Ferley (EDUM); Winnipeg, land, vii (OSUC); Presqu’ile Provincial Park, 15.vii.1932, J.A. Cumming (EDUM); Jobes’ Trail, 10.vii.1996, M. Gurr (DEBU); 11.vii.1960, J.A. Scott (EDUM); 5.viii.1972, S. Presqu’ile Provincial Park, north of day-use 1, Gibb (EDUM). 21.viii.2000, P. Careless (DEBU); Prince Ed- Ontario: Arnprior, 7.vii.1918, 13.vii.1919, ward Co., 15.vii.1914 (2 specimens), Brimley C.M. MacNamara (CNCI); 14.vii.1936, W.J. (ROME); Rockcliffe, 6.vii.1908, E.W. Ingall Brown (CNCI); Avonmore, 13.vii.1939, G.H. (CNCI); 24.vii.1928, J.A. Adams (CNCI); Hammond (CNCI); Beachburg, Renfrew Co., St. Williams, 6.vi.1997, S.A. Marshall (DEBU); 9.vii.1978 (2 specimens), E. Fuller and R. Simcoe, 19.vi.1939, G.E. Shewell (CNCI); Jaagumagi (ROME); Bell’s Corners, South March, 7.vii.1957, S.D. Hicks (CNCI); 18.vii.1948 (8 specimens), S.D. Hicks (CNCI); 24.vi.1959 (MCZC); 30.vi.1959, C.H. Mann 18.vii.1948 (2 specimens), S.D. Hicks (MCZC); (CNCI); Stittsville, 27.vi.1946, Forest Insect Calabogie, Renfrew Co., 26.vi.1978, E. Fuller Survey 046–418 (GLFR); 12.vii.1961, D. (ROME); Chatterton, 19.vii.1951, J.C. Martin, Brown (CNCI); Thousand Islands, Miss Cole- 12.vii.1954, W.A. Dempsey, 13.vii.1954, R.W. man (ROME); Thurlow Township, 18.vii.1940 Smith (CNCI); Constance Bay, Ottawa, (3 specimens) (CNC); Turkey Point, 22.vi.1965, H.F. Howden (CNCI); 24.vi.1965, 17.vii.1940, F.S. Loqier (ROME); Tweed, W.C. McGuffin (LUTB); vii.1970, M. Sanborne 6.vii.1944, G.S. Walley (CNCI); Wagarville, (LEMQ); 11.vii.1976, M. Sanborne (BDMU); 14.vii.1967, H.J. Teskey (MCZC); White Lake, Constance Lake, South March, 20.vi.1939, J.R. 27.vi.1968, J.Robillard (DEBU); ?, 1.vii.1944, Vockeroth (CNCI); Crosby P.O., Kemptville, Forest Insect Survey ONT 1270 (GLFR). 22.vii.1960, Forest Insect Survey 560–447806 Quebec: Abbotsford, 14.vii.1932 (LEMQ); (GLFR); Crotch Lake, 25.vi.1988, J. Denis 22.vi.1937, 25.vi.1937, G.E. Shewell (MCZC); (CNCI); Cummings Bridge, Ottawa, 25.vi.1937 (2 specimens), 26.vi.1937, G.E. 30.vii.1919, J. Thomsen (CNCI); Deep River, Shewell (CNCI); Aylmer, 16.vii.1905, W. 5.vii.1977, D.J. Aspinall (DEBU); Eastview, Metcalfe (CNCI); 15.vi.1913, J.I. Beaulne 1928, R. Painter (CNCI); Foxboro, Sydney Field (CNCI); 30.vi.1919, G.H. Hammond (CNCI); Station, 8.vii.1970, J.F. McAlpine (CNCI); Essex 16.vii.1926, R. Painter (CNCI); 12.vi.1936, Co.,Windsor, Ojibway Prairie, 21.vii.2000, S.A. W.J. Brown (CNCI); Aylmer, Queen’s Park, Marshall (DEBU); 18–19.vi.2002, O. Lonsdale 25.vii.1919, C.B. Hutchings (CNCI); (DEBU); Fuller, Hastings Co., 26.vi.1953 25.viii.1924, A.R. Graham (CNCI); Beech- (ROME); Glenburnie, 2.vii.2001, R. Harmsen Grove, 17.vii.1951, J.F. McAlpine (CNCI); (DEBU); Haldimand-Norfolk Co., Long Point, Bertierville, 16.vii.1950, A. Robert (QMOR); Wilson Tract, 9.vii.1992, R.A. Cannings Deschenes, 23.vi.1930, G.S. Walley (CNCI); (BCPM); Hamilton, 5.vii.1932, B.L. Guyatt 30.vi.1933, R. Ozburn (CNCI); Harrington (ROME); Lambton Co., Walpole Island, Lake, Gatineau Park, 24.vi.1953, S.D. Hicks 4.viii.1996 (12 specimens), J. and A. Ske- (CNCI); Hull, 17.vi.1936, W.J. Brown (CNCI); vington (DEBU, BCPM); La Salle, 20.vii.1966, Iberville Co., Saint-Grégoire, 21.vii.1985, Laro- 21.vii.1967 (2 specimens), K. Stephan chelle and Larivière (LEMQ); Kazabazua, (MCZC); Leonard, 3 mi. E, 12.vii.1970, E.C. 16.viii.1927, G.S. Walley (CNCI); Lac Becker (CNCI); Madoc, 11.vii.1944, G.S. Wal- Roddick, 4.vii.1979, L. Masner (CNCI); Lady- ley (CNCI); Marmora, 23.vi.1942, G.H. Ham- smith, 24.vii.1958, L.A. Kelton (CNCI); La mond (MCZC); 26.vii.1943, 10.vii.1944, G.H. Trappe, 21.vi.1933, 23.vi.1933 (3 specimens), Hammond (CNCI); 25.vi.1952, R. Lambert 25.vi.1933, 30.vi.1933, 4.vii.1933, 17.vii.1934 (CNCI); 2.vii.1957, K.D. Southern (DEBU); (2 specimens), 8.vii.1942, 22.vi.1944, 29.vii.1945, Mer Bleue, 17.vii.1936, W.J. Brown (CNCI); 3.vii.1951, J. Ouellet (QMOR); Laval, Merivale, 21.vi.1943, W.J. Brown (CNCI); Nor- 24.vi.1938 (QMOR); Lévis, T.W. Fyles folk Co., Manestar Tract, 6 km NNW St. Wil- (LEMQ); Masham Township, Gatineau Co., liams, 30.vi.2000, S.A. Marshall (DEBU); 2.vii.1977, O.S. Flint, Jr. (USMN); Montréal, 23.vi.2002, M. Buck (DEBU); N. Burgess vii.1885, G.J. Bowles (LEMQ); 28.vi.1896, Township, Lanark Co., 19.vii.1970, G.C. Wood H.H. Lyman (LEMQ); no date, W. Couper (CNCI); Ottawa, 12.vii.1940, G.A. Hobbs (MCZC); Montréal, Outremont, vi (QMOR); (CNCI); 12.vii.1940, G.A. Hobbs (DEBU); Old Chelsea, King Mountain, 6.viii.1968, J.R. 24.vi.1975, 26.vi.1976, M. Sanborne (BDMU); Vockeroth (CNCI); Otter Lake, 24.vii.1958, L.A.

Kelton (CNCI); Papineau Co, Papineauville, The frons, clypeus, and labrum are yellow or 2.vi.1985, Larochelle and Larivière (LEMQ); brown; a black band crosses the frons below the Pincourt, Île Perrot, 10.vii.1984, V.R. Vickery antennae. Mouthparts are brown, the palps (LEMQ); Rigaud, 8.vii.1920, J. Ouellet brown or yellow. The vertex ranges from or- (QMOR); 24.vi.1939, 29.vi.1939, 3.vii.1939, A. ange to brown and, in British Columbia speci- Robert (QMOR); 26.vi.1941, 27.vi.1942, J. mens, often has a black or brown band across Ouellet (QMOR); 9.vii.1941, A. Robert (QMOR); the posterior. The prothorax is dark brown, often Rigaud, summit of mountain, 7.vii.1986, B.E. yellow anteriorly and posteriorly, with a black Cooper (CNCI); Ste. Annes, 21.vii.1938, band across the apex and base. The front legs 8.vii.1939, D.J. MacDonald (QMOR); are mostly brown, darker on inner faces of the 10.vii.1940, D.J. MacDonald (LEMQ); Saint- femora; the tibiae are sometimes yellow. The Bruno (QMOR); Sainte-Dorothée, 17.vi.1951, middle and hind femora are brown and the tib- 2.vii.1951, 29.vii.1952 (CNCI); Saint-Hilaire, iae and tarsi are yellow; the tibiae have a dark 1.vii.1920, G.A. Moore (LEMQ); Sainte-Rose; mark dorsally. Dorsally, the mesothorax and vii.1896, G.R. Kearley (LEMQ). metathorax have yellow posterior margins. The wings are brown on the anterior half, the colour Other records usually more sharply delineated posteriorly in We have not seen the two Alberta specimens: eastern specimens (Fig. 1). In British Columbia Milk River, Lost River Ranch, vi/vii.1965, D. specimens the brown wash sometimes covers Larson (D. Larson, personal communication); the entire wing, but the pigment is lighter pos- Milk River, 3 km east of North Pinhorn Grazing teriorly. The colour of the abdomen is variable; Reserve Ranch Headquarters, 10.vii.1996, T. often there are large patches of yellow laterally Pike (T. Pike, personal communication). Robert on the basal segments and the abdomen is fre- (1949) recorded other Quebec specimens that quently banded with yellow, black, and brown, we did not examine (he mentioned others noted or more or less mostly reddish brown. Welch above): Montréal, 28.vi.1896, H.H. Lyman; and Kondratieff (1991) described the genitalia, Montréal, vii, A.C. Sheppard; Laval-des- noting that the elongate pseudopenis is distinc- Rapides, 28.vi.1938, G.A. Moore; Fort tive. Coulonge, 17.vii.1919, J.-I. Beaulne; Saint- Hilaire, 11.vii.l938, A.A. Beaulieu; Quebec, Remarks 20.vi.1940, P.E. Mercier. We examined photo- Many authors, at least as early as Wheeler graphs of the following specimens: Kamloops, (1889), have noted the striking coloration of British Columbia, 14.viii.2003, K. West; Perth this insect and have described its mimicry of Road, Ontario, 10.vii.2000, R.-M. Burke. polistine wasps. Some (e.g., Batra 1972) thought that the larval host might be hymen- Description opterous, but Redborg (1998) emphasised that Climaciella brunnea is distinguished by its larvae of C. brunnea, like those of all other robust build, broad frons, and relatively short, mantispines as far as is known, feed on spider thick prothorax (Fig. 1). Measurements of 12 eggs. Wasps are often collected in the vicinity males and 13 females (mean ± SE (except for of congregations of mantispids and, in Utah, ratios) and range in mm): head width, males Batra identified the probable model of the mim- 2.44 ± 0.06 (2.08–2.78), females 2.83 ± 0.13 icry as Polistes fuscatus utahensis Hayward, (2.03–3.63); ratio of frons width to head width, 1933. In Costa Rica, Opler (1981) postulated males 0.41 (0.29–0.44), females 0.42 (0.37– that five separate colour morphs each mimic a 0.48); pronotum length, males 3.62 ± 0.17 different polistine model. Redborg and (2.67–4.40), females 4.23 ± 0.19 (3.00–5.40); MacLeod (1983) cautioned that the limits of ratio of pronotum least width to length, males this variable species are not well known; in Ari- 0.22 (0.17–0.41), females 0.21 (0.18–0.27); zona they reared both yellow-marked and brown forewing length, males 14.27 ± 0.27 (10.65– morphs from the same egg mass. 18.75), females 15.04 ± 0.62 (11.20–19.35); Climaciella brunnea is the most widespread metafemur length, males 3.63 ± 0.19 (2.72– and most frequently collected mantispid in Can- 4.60), females 4.23 ± 0.24 (2.72–5.88). ada. Banks (1911) described C. b. occidentis as Canadian specimens are variably patterned in a western variety. Among other characteristics yellow, brown, and black; yellow areas in many he noted the black vertex band present in many specimens may be darkened to reddish brown. British Columbia specimens. The colour

Fig. 1. Climaciella brunnea. St. Williams, Ontario. Photo: S.A. Marshall.

patterns of eastern and western populations are Quebec). British Columbia dates range from so variable, however, that no subspecific differ- 9 June (Goldstream, Victoria) to 20 October ences hold up under scrutiny. Welch and Kon- (Oyama) (n = 29); eastern ones are from 2 June dratieff (1991) synonymized this variety with (Papineauville, Quebec) to 23 August (Aylmer, the nominate form. Western Canadian speci- Quebec) (n = 121). Eleven dates from British mens (from British Columbia) tend to be larger Columbia are later in the year than the latest than their eastern (Manitoba, Ontario, Quebec) eastern date; seven of these are in September counterparts, however (head width (mm): west- and two are in October. Probably the flight sea- ern males (n = 6), 2.55 ± 0.07 (2.33–2.78); son begins in early June in most years in all eastern males (n = 6), 2.33 ± 0.07 (2.08–2.58); Canadian regions within the range but, at least western females (n = 7), 3.15 ± 0.18 (2.68– in some years, may last 2 months longer in 3.63); eastern females (n = 6), 2.43 ± 0.11 British Columbia than in Manitoba, Ontario, (2.03–2.80)). and Quebec. In body proportions, the most striking differ- Canadian specimens give the following label ences between C. brunnea and the other species data regarding habitat: Douglas-fir trees (Pseu- of Canadian Mantispidae are the broad frons dotsuga menziesii (Mirbel) Franco) (Craig, and the prothoracic proportions. In C. brunnea BC); oak trees (Quercus sp.) (St. Williams, the mean ratio of frons width to head width is ON); ironweed (Vernonia altissima Nutt.) (Wal- 0.41 (n = 25), whereas it is 0.31 (n = 21) pole Island, ON); round-leaved milkweed in L. pulchella and 0.29 (n = 12) in D. inter- (Asclepias ovalifolia Dcne.) in grassland (Milk rupta. The mean of the ratio of prothoracic River, AB); grassland, bare clays with grass, least width to length is 0.21 (n = 25) in cactus (Opuntia polyacantha Haw.), and sage C. brunnea, 0.10 (n = 21) in L. pulchella, 0.14 (Artemisia sp.) (Milk River, AB); grassland ad- (n = 12) in D. sayi, and 0.15 (n = 12) in jacent to open ponderosa pine (Pinus ponderosa D. interrupta. Dougl. ex P. & C. Lawson) woods (Vernon, BC); grassland with bluebunch wheatgrass Natural history (Pseudoroegneria spicata (Pursh) A. Löve) / The flight season in British Columbia differs arrowleaf balsamroot (Balsamorhiza sagittata from that farther east (Manitoba, Ontario, and (Pursh) Nutt.) (Okanagan Falls, BC); at base of

© 2006 Entomological Society of Canada Cannings and Cannings 537 grass clump in Idaho fescue grassland (Festuca Dicromantispa interrupta idahoensis Elmer) (Vernon, BC); three were (Say, 1825), comb. nov. knocked from lakeside willows (Salix sp.) during a rainstorm (Vaseux Lake, BC); one flew Mantispa interrupta Say, 1825; original into a black widow spider (Latrodectus hesperus description; type material, Philadelphia, Chamberlin and Ivie, 1935) web at noon (Oli- Pennsylvania, probably destroyed. ver, BC); and one was collected ambush- Mantispa cincticornis Banks, 1911; original hunting on a dogwood (Cornus sp.) inflores- description; holotype, Brownsville, Texas, MCZC 10768. cence along a forest edge at 1900, in sunshine Mantispa interrupta Say, Welch and Kondra- (Glenburnie, ON). tieff (1991); synonymy of M. cincticornis. Climaciella brunnea is perhaps the most diurnal of North American mantispids and is of- Range ten found in feeding and mating aggregations Restricted in Canada to southern Ontario and on flowers probably formed and maintained Quebec; in the rest of North America, found through the influence of a male pheromone west to Minnesota and south to Florida and (Redborg and MacLeod 1983). southern Mexico (Throne 1972; Hoffman 1992). First-instar larvae spend most of their time standing vertically and motionless on caudal Canadian specimens examined suckers or swaying back and forth with legs Ontario: Chaffey’s Locks, 24.vii.1963, outstretched (Redborg 1998). The larvae must J.C.E. Riotte and I. Smith (ROME); 11.vii.1966 wait for a spider, climb onto it, and reach (2 specimens), Riotte and Kohalmi (ROME); freshly laid eggs before the eggs are encased in 16.vii.1969, 2.viii.1969, 4.viii.1969, J.C.E. silk (Redborg and MacLeod 1983). Unlike Riotte (ROME); 30.viii, 3.x.1969, I.M. Smith D. sayi larvae, those of C. brunnea cannot pen- (ROME); 30.vii.1971, J.C.E. Riotte (ROME); etrate spider egg cases; indeed, C. brunnea lar- 2.vii.1979, F. Phelan (ROME); Chatteron, vae do not even recognize egg cases as food, 7.vii.1951 (CNCI); Hamilton (DEBU); Huntley, and boarding a spider is obligatory for normal 19.vii.1960, D. Brown (CNCI); Lanark, Pat- development (Redborg and MacLeod 1983). terson Lake, 13.viii.1977, S.A. Marshall Larvae sit on the dorsum of the carapace with (DEBU); Marmora, 15.vii.1951, R. Lambert their mouthparts adjacent to the membrane be- (CNCI); Niagara-on-the-Lake, 14.viii.1932, G.G. Duston (DEBU); North Burgess Township, tween the carapace and the leg bases (Redborg Lanark Co., 27.vii.1967, 18.ix.1967, D.M. 1998). Wood (CNCI), 21.vi.1970, 5.vii.1970, G.C. First-instar larvae overwinter on spiders, at Wood (CNCI); Ottawa–Carleton Regional Mu- least in the north. Redborg and MacLeod (1983) nicipality, Granite Hill farm, 1.ix.1981 (BMNH); discussed this phoretic behaviour and develop- Perth Road, Frontenac Co., 28.vii.1971, P. Ward ment in detail. Host range is rather narrow: all and J. Edsall (CNCI); 23.vii.1971 (2 speci- documented wild-caught hosts are lycosid spi- mens), U. Luciuk, 15.viii.1970, P. Ward and J. ders from the genera Schizocosa, Lycosa (Red- Edsall (ROME); Stirling, 25.viii.1963, A.F. borg and MacLeod 1983; LaSalle 1986), Johnson (DEBU); Walsh, 28.vi.1915 (DEBU). Rabidosa (Redborg 1998), and Tarantula Quebec: Aylmer, Queen’s Park, 29.vii.1946, (George and George 1975). Larvae that board G.S. Walley (CNCI); Burnet, 12.vii.1966, Mar- male spiders can switch to females during spi- guerite Job (CNCI); Kirk’s-Ferry, 9.viii.1950, B.P. Beirne (CNCI); La Trappe, 8.vii.1942, der mating or cannibalism of males by females 22.vi.1944, 3.vii.1951, J. Ouellet (QMOR); (Scheffer 1992). Redborg and Redborg (2000) Luskville, Belisle Beach, 12–19.viii.1961, E.G. found C. brunnea on 19% of Schizocosa sp. in Munroe (CNCI); Pontiac, Eardley, 26.vi.1991, forest leaf litter, while sympatric D. sayi (= S. LaPlante (CNCI); Rigaud, 3.vii.1939, A. Rob- Mantispa uhleri Banks) were phoretic on spider ert (QMOR). species above the forest floor. On an adjacent Robert (1949) listed the Aylmer, La Trappe grassy field, the two mantispids were both (1942, 1944), and Rigaud records noted above. found on the lycosid Rabidosa punctulata Riotte (1968) commented on the Chaffey’s Locks, (Walckenaer, 1837). Ontario, collections and Barr and Barr (1980)

© 2006 Entomological Society of Canada 538 Can. Entomol. Vol. 138, 2006 published a photograph of a specimen from the Lake, one was “taken sweeping edge of oak same locality. bush” (label data). Hoffman and Brushwein (1990) and Redborg and MacLeod (1985) suggested that D. inter- Description rupta larvae board spiders and enter spider egg Dicromantispa interrupta is the largest spe- sacs as they are produced, although Redborg cies of Canadian Mantispidae, at least in most (1998) indicated that they may also be able to linear measurements. Measurements of 6 males penetrate completed egg sacs. In South Caro- and 6 females (mean ± SE and range in mm): lina, Hoffman and Brushwein (1990) recorded head width, males 2.56 ± 0.13 (1.95–2.85), fe- immature stages of D. interrupta associated males 2.77 ± 0.15 (2.23–3.25); ratio of frons with 10 species of spiders from the Gnapho- width to head width, males 0.31 (0.28–0.32), sidae, Lycosidae, Ctenidae, and Pisauridae. females 0.31 (0.29–0.34); pronotum length, Host species are predominantly ground- males 4.77 ± 0.23 (3.70–5.28), females 5.29 ± wandering hunters. Having boarded the host, 0.21 (4.46–5.92); ratio of pronotum least width the larva wraps tightly around the spider’s pedi- to length, males 0.15 (0.13–0.16), females 0.14 cel (Redborg 1998); during a spider molt, the (0.10–0.17); forewing length, males 16.4 ± 0.85 mantispid may enter the host’s book lungs (12.40–18.30), females 18.33 ± 0.97 (15.00– (Hoffman and Brushwein 1990; Redborg 1998) 21.75); metafemur length, males 3.8 ± 0.24 if the opening is large enough. (2.75–4.44), females 4.19 ± 0.20 (3.58–4.84). Specimens are mainly yellow with variable Dicromantispa sayi (Banks, 1897) brown markings. The frons has a brown mid- line, usually broader between the antennae and Mantispa sayi Banks, 1897; original description; usually widening below on the clypeus and lectotype male, Brazos Co., Texas, MCZC labrum. The vertex behind the antennae some- 10767. times has brown markings and the whole head Mantispa fuscicornis Banks, 1911; original is washed with brown in some specimens. The description; lectotype male, Kissimmee, mouthparts are usually brown; the antennae are Florida, MCZC 10769. yellow. The prothorax has a middorsal brown Mantispa uhleri Banks, 1943; original stripe at the anterior and posterior ends; the in- description; holotype, Pennsylvania, MCZC ner faces of the profemora and protibiae are 22132. brown. The wings are clear with mostly yellow Mantispa sayi Banks, Hoffman (1989); synonymy venation and are distinctively marked with of M. uhleri and M. fuscicornis. brown in the costal/radial space (especially in Dicromantispa sayi (Banks), Hoffman (2002); the pterostigma area), at the origins of the veins new genus. along the radius, at the wing tips and, to a lesser extent, at the wing bases. The brown Range markings on the abdomen are variable, but usu- In Canada the species is known only from ally a dark middorsal stripe occurs. Welch and extreme southern Ontario from the Windsor Kondratieff (1991) described the male and fe- area east to Niagara-on-the-Lake. In the United male genitalia, noting that the epiproct and other States, it occurs across most of the eastern states male terminalia structures are distinctive. south to Florida and west to South Dakota, Ne- braska, Utah, and eastern Arizona. It ranges southwards through Mexico to Panama and is Natural history also found in the Bahamas, Cuba, and Puerto The flight season, based on Canadian records, Rico. is 21 June (North Burgess Township, Lanark Co., Ontario) to 3 October (Chaffey’s Locks, Canadian specimens examined Ontario). Recorded sites of capture are usually Ontario: Essex Co., Point Pelee National trees and shrubs, although one specimen Park, West Beach, 20.vii.2000, S.A. Marshall (Lanark Co., Ontario, 13.viii.1997) was col- (DEBU); Point Pelee National Park, group lected at lights at night. Throne (1972) reported campground, mercury vapour (mv) light, sweeping specimens from oak, eastern white 24.vii.2003, S.M. Paiero (DEBU); Point Pelee pine (Pinus strobus L.), and eastern redcedar National Park, Northwest Beach, mv light, (Juniperus virginiana L.). At Niagara-on-the- 29.vii.2003, D. Cheung (DEBU); Essex Co.,

Windsor, Ojibway Prairie, 4.ix.1994. W. Bennett present ventrally on the femora and dark marks (DEBU); Harrow, 17.vii.1976, C.D. Neilson on the trochanters. The meso- and meta-coxae (DEBU); Kent Co., Rondeau Provincial Park, are largely brown in females but the latter are 28.vii.1967, I.M. Smith (ROME); Rondeau Pro- mostly pale in males; in both sexes the poste- vincial Park, Marsh Trail North, 42°18′N, rior faces of the metacoxae are brown or spot- 81°51′W, 10.viii.2003, M. Buck (DEBU); ted with brown. The pleural sclerites are mostly 20.vii.2004, S.M. Paiero (DEBU); Rondeau brown and margined with pale lines, but those Provincial Park, Visitor Centre, 42°46′50′′N, of the metathorax in males are pale. The meso- 81°50′38′′W, mv light, 14.viii.2003, S.M. thorax and metathorax are largely brown dor- Paiero (DEBU); Rondeau Provincial Park, sally but the dark areas on the mesothorax are group campground, 42°17′35′′N, 81°50′52′′W, normally divided into lateral and medial 20–22.vii.2004 (3 specimens), S.A. Marshall patches. The wings are clear with brown vena- (DEBU), (2 specimens) S.M. Paiero (DEBU); tion; other brown pigment is mainly restricted Niagara-on-the-Lake, 15.vii.1934, W.L. Putman to the costal/radial space (pterostigma) midway (DEBU); Norfolk Co., Turkey Point Tract, along the wing. The abdomen is strongly marked 18.vii.2001, A. Timpf and M. Gartshore (DEBU); with black or brown on the sternites and mid- Wheatley, 22.vii.1976, M.J. Sharkey (DEBU). dorsally and posterolaterally on the tergites. As indicated above, females usually are more ex- Description tensively darkened on the thorax than males; Measurements of 6 males and 6 females this is also true of the abdomen, which some- (mean ± SE and range in mm): head width, times is all black. Specimens from the southern males 2.14 ± 0.15 (1.58–2.70), females 2.20 ± parts of the range in the United States (formerly 0.18 (1.58–2.75); ratio of frons width to head known as M. fuscicornis) are darker overall; es- width, males 0.29 (0.27–0.30), females 0.29 pecially notable is the pale brown wash on the (0.28–0.30); pronotum length, males 3.78 ± wing membrane posterior to the first radial vein 0.26 (2.88–4.80), females 4.21 ± 0.31 (2.60– in some specimens. Welch and Kondratieff 4.84); ratio of pronotum least width to length, (1991) figured the male and female genitalia males 0.14 (0.10–0.17), females 0.14 (0.13– and described the male’s distinctive epiprocts 0.15); forewing length, males 12.10 ± 0.88 with their spine-bearing projections. (9.45–15.45), females 14.05 ± 0.94 (9.75– 16.95); metafemur length, males 2.90 ± 0.24 Remarks (2.08–3.92), females 3.14 ± 0.23 (2.00–3.80). If we assume D. sayi is a single species, it is The head, body, and legs are pale yellow with extremely variable in colour over its large range. extremely variable brown or black markings; Indeed, three species were originally named to Figure 2 shows one of the darkest specimens. account for this variation: M. fuscicornis from Canadian specimens usually have the face with Florida and Texas, M. sayi mostly west of the a dark median vertical stripe from the labrum to Mississippi River and in Florida, and M. uhleri the antennae, the stripe branching in a Y-shaped east of the Mississippi and northward. MacLeod or trident-shaped mark behind the antennae and (in Hughes-Schrader 1979) and Redborg (1982a) forming an extensive pattern of dark lines on considered them sibling species; Hughes- the vertex, usually isolating several pale spots — Schrader (1979) found no cytological differ- a pair immediately behind the antennae and ences among them. Hoffman (1989; personal three larger ones (the middle one more elon- communication) synonymized them under gate) crossing the posterior half of the vertex. M. sayi, finding absolutely no genitalic differ- In some specimens the dark pigment spreads ences among populations from the northeastern over much of the vertex, obscuring these spots. United States to Panama. Nevertheless, Redborg The mandibles are brown; the antennae are brown (1998; personal communication), based on his with the basal segments yellow. The prothorax field experience with live insects, maintains that ranges from mostly yellow to mostly brown, but these taxa should still be considered separate the anterolateral corners are dark and usually a until more evidence of their status is presented. middorsal stripe is visible, often present only He has collected and reared hundreds of speci- anteriorly and posteriorly. The procoxae are mens of the darker, more eastern (uhleri) form normally lined with brown ventrally, and the (with little or no colour variation) from wooded profemora and protibiae are mostly brown. The areas in eastern Iowa, Illinois, Indiana, and other legs are mostly pale, with dark lines often Minnesota, evidently representing the western

Fig. 2. Dicromantispa sayi. Point Pelee, Ontario. Photo: S.A. Marshall.

edges of the historic eastern deciduous forest. Ontario, while a darker one occurs in Carolin- He considers the paler (sayi) form a sibling spe- ian woodland sites elsewhere along the Lake cies of the historic grassland areas stretching Erie shore (Paiero et al. 2006; S. Marshall, per- west from Iowa; he has also collected adults in sonal communication). However, we can find sandy areas of central Illinois that support pines no real evidence in the small sample that there and live oaks (K. Redborg, personal communi- are two separate populations, one dark and one cation). pale, that equate to the original species, Nevertheless, we find Hoffman’s taxonomic D. uhleri and D. sayi, respectively. Until more work compelling and have followed his system- taxonomic, genetic, molecular, or ecological atic revisions, calling the combined species work clarifies this situation, we prefer to con- D. sayi, but we are sympathetic to Redborg’s sider all known Canadian specimens as consti- conservative position on the matter. If biological information is recorded under only one name tuting one species, D. sayi. This species has not and the three taxa, after further study, are con- been previously reported in the literature from firmed as different, it will be difficult to sort Canada, although Kevan (1979) suspected that out the details, at least in regions where the it occurred in the country. populations are sympatric. The most northerly American populations for- Natural history merly were identified as Mantispa (= Dicro- The extensive work of Redborg (1982a, mantispa) uhleri, and most Canadian specimens have the dark coloration characteristic of this 1982b, 1983) and Redborg and MacLeod taxon (Fig. 2). Four or five specimens with a (1985) on the developmental ecology of this similar, but somewhat lighter, colour pattern species (cited as M. uhleri) has revealed more come from scattered localities in the Canadian about its biology than is known for any other range, including the remnant tallgrass prairie at North American mantispid. In Canada all re- Ojibway Prairie, Windsor. This has suggested cords are from the Carolinian zone near Lake that a pale-coloured population (i.e., the origi- Erie; dates are from 15 July to 4 September nal D. sayi) might be typical of remnant tall- (n = 17). Redborg and MacLeod (1985) light- grass prairie habitats in extreme southwestern trapped adults in Illinois from mid-June to mid-

October, with the majority caught from July Canadian specimens examined through mid-September. British Columbia: Oliver, 12.vii.1985, M. Most first-instar larvae probably are spider Sarell (UVIC); Oliver, Gallagher Lake, boarders, entering the egg sac as it is spun by 13.viii.1985, M. Sarell (BCPM); Oliver, UBC the host; however, they can also find and pene- Geology Camp, 19.vii.1989, 22.vii.1990, trate sacs already constructed (Redborg and 23.vii.1990, S.G. Cannings (BCPM, SMDV); MacLeod 1985). Larvae usually fix themselves Osoyoos, East Bench, 6.vi.2001, J. Scudder to the spider’s pedicel and then enter the host’s (SMDV), 23.vii.2001, G. Scudder (SMDV); book lungs (Redborg 1998), where they feed on Summerland, Trout Creek Point, 16.vii.1977, L. haemolymph; such parasitism increases the de- Dale (BCPM); Vaseux Lake, 26.vi.1959, L.A. velopment time and decreases the adult size of Kelton (CNCI); Vernon, 21.viii.1924, E.A. the spider (Redborg 1982b). Larvae can switch Rendell (CNCI). hosts during spider cannibalism (O’Brien and Ontario: Windsor, Ojibway Prairie, Redborg 1997); this is perhaps more efficient 13.viii.2002, S.A. Marshall (DEBU). than switching during copulation, given the short copulation times of the major host, Philo- Description dromus vulgaris (Hentz, 1847) (Redborg and Leptomantispa pulchella is the smallest of MacLeod 1985). A wide variety of hunting spi- the Canadian mantispids. Measurements of 11 ders are boarded: 31 species in 21 genera from males and 10 females: head width, males 1.83 ± almost all families of Lycosoidea and Club- 0.06 (1.38–2.00), females 1.86 ± 0.09 (1.35– ionoidea have been documented (Redborg and 2.25); ratio of frons width to head width, males MacLeod 1985). The first-instar larva over- 0.30 (0.28–0.32), females 0.32 (0.30–0.36); winters on the spider host and, in Illinois, the pronotum length, males 3.26 ± 0.08 (2.64– first adult mantispids emerge in late June from 3.60), females 3.15 ± 0.14 (2.32–3.80); ratio of late-spring or early-summer egg sacs of spiders pronotum least width to length, males 0.10 that overwintered as adults. From one to three (0.08–0.12), females 0.22 (0.09–0.12); forewing generations of mantispids could be produced by length, males 9.44 ± 0.23 (7.20–11.75), females any given lineage each year, depending on the 9.44 ± 0.50 (6.60–11.55); metafemur length, life history of the spider (Redborg and Mac- males 2.14 ± 0.07 (1.64–2.44), females 2.13 ± Leod 1985). 0.12 (1.48–2.60). Canadian specimens are yellow, patterned in brown and black (Fig. 3). The head is yellow with a black vertical stripe on the face forking Leptomantispa pulchella between the antennal bases; there is a small (Banks, 1912) brown spot between the arms of the fork at the front of the vertex. Although any brown marks Mantispilla pulchella Banks, 1912; original on the vertex are frequently diffuse, there is of- description; holotype, Eureka, Utah, MCZC ten a brown bar, sometimes interrupted, cross- 10766. ing the centre of the vertex from eye to eye. Mantispa pulchella Nakahara, 1913; synonymy. The antennae are brown or black with the basal Leptomantispa pulchella (Banks), Hoffman segment yellow. The prothorax is mostly (2002); synonymy. brown, although dorsally it is yellow, except for the base. The legs are mostly yellow; the pro- femur has a brown inner face and marks at the Range apex and base of the outer face; the mesofemur In Canada L. pulchella is known from the and metafemur bear a brown line ventrally in Okanagan Valley in south-central British Co- the basal half. The meso- and meta-coxae are lumbia and the Windsor area of extreme marked with brown. The mesothorax and meta- southwestern Ontario. In the United States it thorax each have a pair of wide brown stripes ranges over most of the western states, south dorsally; the lateral thoracic sclerites and coxae through California and Utah to Arizona and are margined and spotted with brown. The wings Texas, east to Illinois, Ohio, and New York, and are clear; the venation ranges from yellow to south to the Carolinas and Georgia. Farther dark brown; the radius is yellow or light brown; south, L. pulchella ranges from Mexico to Costa the pterostigma is orange. The venter and dorsal Rica and is also recorded in Cuba. margins of the abdominal sternites are dark

Fig. 3. Leptomantispa pulchella. Ojibway Prairie, Windsor, Ontario. Photo: S.A. Marshall.

brown or black; the tergites have paired brown obvious. Leptomantispa pulchella has not been spots dorsally and vertically paired stripes later- previously reported from Canada. ally. Welch and Kondratieff (1991) showed that the structure of the epiproct is different from Natural history that of the two Canadian Dicromantispa spe- In the Okanagan Valley, L. pulchella appar- cies; the ventromedial lobes are reduced to a ently is both diurnal and nocturnal. During the patch of spines. day, specimens have been taken from the side of a tent, from vegetation, and from a Malaise trap in open ponderosa pine woods. They have Remarks also been captured in houses and at night in a The size of British Columbia specimens is, mercury vapour lamp in pine woods. The cap- on average, greater than that of the United States tures range from 6 June to 21 August (n = 10). specimens examined. For example, in males, The Ontario specimen was collected on 13 Au- the head width of BC specimens (n =5)is gust at Ojibway Prairie, a relict tallgrass prairie 1.96 ± 0.02 mm (1.18–2.00 mm), while that of at Windsor. US specimens (n = 6) is 1.72 ± 0.08 mm (1.38– The developmental ecology of the species has 1.90 mm). The forewing length is 10.02 ± been detailed by Hoffman and Brushwein 0.35 mm (9.30–11.75 mm) in BC material and (1989). First-instar larvae board spiders to gain access to freshly laid eggs; they will not feed on 8.96 ± 0.43 mm (7.20–9.90 mm) in US mate- eggs unless they have spent some time aboard rial. The means of these two measurements spiders. Their obligatory boarding behaviour is from the single Ontario male (1.49 mm and more similar to that of C. brunnea larvae than 8.00 mm, respectively) are similar to those of to that of other Nearctic mantispid larvae. smaller US specimens. The Ontario specimen is These larvae are usually found singly on the less strongly marked with dark pigment than dorsum of the spider pedicel, where apparently the BC specimens. Most specimens we have they feed on the host’s haemolymph. The spi- examined from the eastern United States have ders involved are all commonly found on fo- the yellow base colour darkened to light brown, liage: Anyphaenidae, 65.7%; Salticidae, 19.4%; rendering the brown colour pattern less Clubionidae, 10.4%; and others, 4.5%. These

© 2006 Entomological Society of Canada Cannings and Cannings 543 spiders also build silken retreats on leaves and Banks, N. 1911. Descriptions of new species of North under bark; they may be easier for mantispid American neuropteroid insects. Transactions of the larvae to find and board than other species that American Entomological Society, 37: 335–360. do not use such retreats. Banks, N. 1912. Notes on Nearctic Mantispidae. Pro- ceedings of the Entomological Society of Washing- ton, 14: 178–179. Banks, N. 1943. New Neuroptera and Trichoptera Acknowledgements from the United States. Psyche (Cambridge), 50: 74–81. We thank the following people, who made Barr, D., and Barr, N. 1980. The miniature world of material available for study: P. Arnaud, Jr. (Cal- Aubrey Critch. Readers Digest, 117: 178–183. ifornia Academy of Sciences, San Francisco, Batra, S.W.T. 1972. Notes on the behavior and ecol- California), R. Duncan (Pacific Forestry Centre, ogy of the mantispid Climaciella brunnea occi- Victoria, British Columbia), N. Duffey (Lyman dentalis [sic]. Journal of the Kansas Entomological Entomological Museum, McGill University, Society, 45: 334–340. Sainte-Anne-de-Bellevue, Quebec), O. Flint, Jr. Davidson, J.A. 1969. Rearing Mantispa viridis Walker (US National Museum, Washington, D.C.), T. in the laboratory (Neuroptera, Mantispidae). Ento- Galloway (University of Manitoba, Winnipeg, mological News, 80: 29–31. Manitoba), K. Kevan (Lyman Entomological Enderlein, G. 1910. Klassifikation der Mantispiden Museum), S. Marshall (University of Guelph, nach dem material des Stettiner Zoologischen Museums. Stettiner Entomologische Zeitung, 71: Guelph, Ontario), J. Martin (CNCI, Agriculture 341–379. Canada, Ottawa, Ontario), N. Penny (California George, L.D., and George, N.L. 1975. Notes on the Academy of Sciences), M. Thayer (Museum of three hundred and fifty-eighth meeting: a new re- Comparative Zoology, Harvard University, cord of mantispid reared from spider. Pan-Pacific Cambridge, Massachusetts), P. Harper, (Uni- Entomologist, 51: 90. versité de Montréal, Montréal, Quebec), G. Hoffman, K.M. 1989. Taxonomic status of Mantispa Scudder (Spencer Entomological Museum, Uni- sayi, Mantispa fuscicornis and Mantispa uhleri versity of British Columbia, Vancouver, British (Neuroptera: Mantispidae). Proceedings of the En- Columbia), and G. Wiggins (Royal Ontario Mu- tomological Society of Washington, 91: 637–639. seum, Toronto, Ontario). P. Belton, R.-M. Hoffman, K.M. 1992. Systematics of the Burke, S. Ford, K. Hoffman, K. Iverson, D. Mantispinae (Neuroptera: Mantispidae) of North, Central and South America. Ph.D. thesis, Larson, K. MacKenzie, D. McMullen, K. Need- Clemson University, Clemson, South Carolina. ham, D. Nicholson, S. Paiero, H. Philip, T. Hoffman, K.M. 2002. Family Mantispidae. In A guide Pike, K. Redborg, M. Sarell, J. Skevington, G. to the lacewings (Neuroptera) of Costa Rica. Scudder, K. West, T. Wheeler, and M. Wood Edited by N. Penny. Proceedings of the California also supplied specimens and information. C. Academy of Sciences, 53: 161–457. pp. 251–275. Borkent helped with the production of the Hoffman, K.M., and Brushwein, J.R. 1989. Species manuscript. K. Hoffman, S. Marshall, and K. of spiders (Araneae) associated with the immature Redborg commented on manuscript drafts; we stages of Mantispa pulchella (Neuroptera, Man- particularly thank them for their help. K. tispidae). Journal of Arachnology, 17: 7–14. Hoffman generously provided unpublished in- Hoffman, K.M., and Brushwein, J.R. 1990. Spider formation on taxonomic matters and suggested (Araneae) taxa associated with the immature stages of Mantispa interrupta (Neuroptera: Man- that I publish the new combination of Dicro- tispidae). Entomological News, 101: 23–28. mantispa interrupta (Say). S. Marshall kindly Hughes-Schrader, S. 1979. Diversity of chromo- allowed us to reproduce the photographs in Fig- somal segregational mechanisms in mantispids ures 1–3. (Neuroptera: Mantispidae). Chromosoma, 75:1– 17. Imms, A.D. 1957. A general textbook of entomol- References ogy. Methuen, London. Kevan, D.K.McE. 1979. Neuroptera. In Canada and Arnett, R.H., Jr., Samuelson, G.A., and Nishida, G.M. its insect fauna. Edited by H.V. Danks. Memoirs 1993. The insect and spider collections of the of the Entomological Society of Canada, 108:1– world. 2nd ed. Sandhill Crane Press, Gainesville, 573. pp. 354–356. Florida. LaSalle, M.W. 1986. Notes on the mantispid Clima- Banks, N. 1897. New North American neuropteroid ciella brunnea (Neuroptera: Mantispidae) in a insects. Transactions of the American Entomolog- coastal marsh habitat. Entomological News,97: ical Society, 24: 21–31. 7–10.

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